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PHILADELPHIA - When the University of Pennsylvania kicked off to Dartmouth to start Saturday's football game at Franklin Field, junior Kyle Wilcox was nowhere to be found among the running backs on Penn's depth chart.

Three hours and 49 minutes later -- after the Quakers and the Big Green had played 60 minutes of regulation football, three full overtimes, and one possession of a fourth -- it was Wilcox who dramatically ended things.

With Penn facing a third-and-5 at the 20-yard line and needing just a field goal to win this marathon, Wilcox took a handoff, bounced around the right side, stiff-armed a Big Green defender, and beat the rest of them to the sideline. The next thing you know, he was in the end zone and Penn had escaped with a 37-31 victory.

It ended the longest game in Ivy League history, one that featured blocked field goals, fumbles returned for touchdowns, and back-and-forth action that left fans on both sides completely drained by the end.

It was also the Ivy opener for both teams as Penn improved to 2-1 on the year and 1-0 in league play with its sixth straight win over the Big Green. Dartmouth, which has still not won in Philadelphia since 1997, dropped to 1-2 this season and 0-1 in Ivy play.

Wilcox had the winner, but the biggest hero of the day was senior linebacker David Park. He was all over the place for the Quakers on Saturday, with two of the biggest plays of the game coming via his hands. In the second quarter, he gave Penn a 14-7 lead with an 84-yard fumble recovery for a touchdown.

While that play was huge, the one he made at the end of regulation was gigantic.

With just four seconds left in regulation, and the game tied at 21-21, Dartmouth lined up for the game-winning 21-yard field goal. However, Park again stepped up, blocking Riley Lyons' chip shot and stunningly sending the game to overtime.

Penn had the first possession in overtime, and drove all the way to the 5-yard line. However, on fourth-and-1 the Quakers opted to go for it and instead were stuffed at the line of scrimmage.

Dartmouth suddenly had the game in its hands again. But Penn quickly forced a fourth down and Lyons -- maybe still thinking about his kick in regulation -- hooked his 34-yard attempt wide left to give the Quakers new life once again.

Dartmouth started the second OT with the ball, and needed just three plays to hit paydirt. Quarterback Dalyn Williams hit Dominick Pierre for 12 yards on the third play of the drive, then Kirby Schoenthaler in the end zone on the next play.

The Quakers responded, led by QB Billy Ragone. On a day when he became just the second player in Penn history to go over 6,000 career yards in total offense, it was Ragone who got Penn to the third overtime. Facing third-and-12 at the 27-yard line, he escaped a collapsing pocket and found daylight along the Penn sideline where he rumbled all the way to the end zone. That tied the score at 28-28 and sent the game into a third overtime.

Penn started with the ball, and after three plays the Quakers put their faith in junior Connor Loftus. He rewarded them by putting his 38-yard field goal between the uprights.

Needing a field goal to tie -- and knowing a touchdown would win it -- Dartmouth went nowhere. Williams sandwiched a 1-yard run between a pair of incomplete passes, putting the game back on the foot of Lyons. This time he delivered, knocking home a 40-yard field goal to knot the game at 31-31.

Dartmouth started the fourth OT with the ball, and appeared to have a touchdown when Williams found Pierre in the end zone. However, the Big Green were called for a personal foul which nullified the play. After that, Dartmouth got no closer than the 20-yard line, and on fourth down Lyons was called upon to kick a 42-yard field goal. Wide left, again.

That set the stage for Penn's possession, and after Wilcox was stopped after two yards on the left side, and three yards up the middle, the right side proved to be the right choice.

Nearly four hours earlier, Dartmouth struck first on Saturday, getting on the board thanks to a one-yard run from Pierre, but the defending Ivy League champions answered in a hurry, primarily on the arm of Ragone. His go-to receiver early in the drive was senior Ryan Mitchell, whom he found for a 14-yard completion early on a key third-down conversion. Later, the duo connected on a 52-yard pass that moved the ball all the way down to the Dartmouth 8-yard line.

Just as the Dartmouth defense looked as though it might hold the Quakers to a field goal, Ragone capped the drive by scrambling to his right and finding senior wideout Ty Taylor in the corner of the end zone for the touchdown. That tied the game at 7-7 at the 5:12 mark of the opening quarter.

Things hit a bit of a lull at the end of the first quarter and spanning past the halfway point of the second quarter, with the two teams combining for four punts and no scores.

Dartmouth looked to end the streak of punts after Williams connected with Victor Williams for a 48-yard pass to move the ball to the Penn 28. On third down from the 16-yard line, a ferocious rush from Penn's front seven allowed senior Sam Chwarzynski an open shot on Williams, forcing a fumble that Park scooped up and took 84 yards for the touchdown.

The Quaker defense wasn’t done yet, though. The Big Green had a chance to even things up at 14 with under two minutes left in the half, but safety Evan Jackson swatted down a fourth-and-2 pass that would have moved Dartmouth inside the red zone.

That was all the action in the first half, as the Quakers led at the break, 14-7.

Dartmouth tied the game early in the third quarter, taking advantage of an interception to drive 44 yards in just six plays. Williams found Dean Bakes over the middle for 11 yards and the touchdown.

Penn answered with a classic Quaker drive. Running 14 plays and taking nearly eight minutes, the Red and Blue chipped away at the Dartmouth defense and split its ground game and air attack nicely. The Quakers regained the lead when sophomore Mike Elespuru rushed it in from three yards out with 1:37 left in the third. Ragone was again superb on the drive, going 5-of-5 passing for 63 yards and running for a big first down late in the drive.

Dartmouth tied it again with 10:18 to play in the fourth. Starting on its own 31, the Big Green got a huge lift when Pierre broke away for 58 yards. He was taken down by Evan Jackson at the 2-yard line, but that only delayed the inevitable as Williams ran it in on the next play.

Penn ran 81 plays and finished with 449 yards of total offense compared to Dartmouth's 534 yards on 91 plays. Most importantly, Penn was 4-for-5 in the red zone, while holding Dartmouth to just four scores in eight red zone trips.

Wilcox ran for 94 yards on 18 carries, most of which came late in the fourth quarter and overtime. Ragone was 17-of-28 for 213 yards and a touchdown through the air and added 60 yards and a score on the ground. Senior Ryan Mitchell had his first career 100-yard receiving game with six catches for 100 yards. Taylor caught five passes for 39 yards and his second touchdown of the year, while sophomore Cameron Countryman added three catches for a career-best 52 yards.

Dartmouth had not allowed a sack this season entering the game, but the Quakers finished with four. In addition to Park and Chwarzysnki, Tyler Drake and Tanner Thexton each added their first career sack. Seniors Dan Wilk and Sebastian Jaskowski had a team-high eight tackles, as did junior linebacker Dan Davis. Junior Brion Wood added a key interception late in the fourth quarter to halt a drive deep in Penn territory.

Penn wraps up its non-conference slate next Saturday, taking on William & Mary at the Tribe's Zable Stadium. It will mark Penn's first game in Williamsburg, Va., since 1995. Kickoff is set for 3:30 p.m.